Light quality is a crucial physical factor driving coral distribution along depth gradients. Currently, a 30 m depth limit, based on SCUBA regulations, separates shallow and deep mesophotic coral ecosystems (MCEs). This definition, however, fails to explicitly accommodate environmental variation. Here, we posit a novel definition for a regional or reef‐to‐reef outlook of MCEs based on the light vs. coral community–structure relationship. A combination of physical and ecological methods enabled us to clarify the ambiguity in relation to the mesophotic definition. To characterize coral community structure with respect to the light environment, we conducted wide‐scale spatial studies at five sites along shallow and MCEs of the Gulf of Eilat/Aqaba (0–100 m depth). Surveys were conducted by technical‐diving and drop‐cameras, in addition to one year of light spectral measurements. We quantify two distinct coral assemblages: shallow (<40 m) and MCEs (40–100 m), exhibiting markedly different relationships with light. The depth ranges and morphology of 47 coral genera were better explained by light than depth, mainly, due to photosynthetically active radiation (PAR) and ultraviolet radiation (UVR) (1% at 76 and 36 m, respectively). Branching coral species were found mainly at shallower depths, that is, down to 36 m. Among the abundant upper‐mesophotic specialist corals, Leptoseris glabra, Euphyllia paradivisa, and Alveopora spp. were found strictly between 40 and 80 m depth. The only lower‐mesophotic specialist, Leptoseris fragilis, was found deeper than 80 m. We suggest that shallow coral genera are light‐limited below a level of 1.25% surface PAR and that the optimal PAR for mesophotic communities is at 7.5%. This study contributes to moving MCE ecology from a descriptive phase into identifying key ecological and physiological processes structuring MCE coral communities. Moreover, it may serve as a model enabling the description of a coral zonation worldwide on the basis of light quality data.

Aim Mesophotic coral ecosystems (MCEs) are unique communities that support a high proportion of depth‐endemic species distinct from shallow‐water coral reefs. However, there is currently little consensus on the boundaries between shallow and mesophotic coral reefs and between upper versus lower MCEs because studies of these communities are often site specific. Here, we examine the ecological evidence for community breaks, defined here as species loss, in fish and benthic taxa between shallow reefs and MCEs globally. Location Global MCEs. Time period 1973–2017. Major taxa studied Macrophytes, Porifera, Scleractinia, Hydrozoa, Octocorallia, Antipatharia and teleost fishes. Methods We used random‐effects models and breakpoint analyses on presence/absence data to identify regions of higher than expected species loss along a depth gradient of 1–69 m, based on a meta‐analysis of 26 studies spanning diverse photoautotrophic and heterotrophic taxa. We then investigated the extent to which points of high faunal turnover can be explained by environmental factors, including light, temperature and nutrient availability. Results We found evidence for a community break, indicated by a significant loss of shallow‐water taxa, at ~ 60 m across several taxonomically and functionally diverse benthic groups and geographical regions. The breakpoint in benthic composition is best explained by decreasing light, which is correlated with the optical depths between 10 and 1% of surface irradiance. A concurrent shift in the availability of nutrients, both dissolved and particulate organic matter, and a shift from photoautotroph to heterotroph‐dominated assemblages also occurs at ~ 60 m depth. Main conclusions We found evidence for global community breaks across multiple benthic taxa at ~ 60 m depth, indicative of distinct community transitions between shallow and mesophotic coral ecosystems. Changes in the underwater light environment and the availability of trophic resources along the depth gradient are the most parsimonious explanations for the observed patterns.

Shallow water zooxanthellate coral reefs grade into ecologically distinct mesophotic coral ecosystems (MCEs) deeper in the euphotic zone. MCEs are widely considered to start at an absolute depth limit of 30m deep, possibly failing to recognise that these are distinct ecological communities that may shift shallower or deeper depending on local environmental conditions. This study aimed to explore whether MCEs represent distinct biological communities, the upper boundary of which can be defined and whether the depth at which they occur may vary above or below 30m. Mixed-gas diving and closed-circuit rebreathers were used to quantitatively survey benthic communities across shallow to mesophotic reef gradients around the island of Utila, Honduras. Depths of up to 85m were sampled, covering the vertical range of the zooxanthellate corals around Utila. We investigate vertical reef zonation using a variety of ecological metrics to identify community shifts with depth, and the appropriateness of different metrics to define the upper MCE boundary. Patterns observed in scleractinian community composition varied between ordination analyses and approaches utilising biodiversity indices. Indices and richness approaches revealed vertical community transition was a gradation. Ordination approaches suggest the possibility of recognising two scleractinian assemblages. We could detect a mesophotic and shallow community while illustrating that belief in a static depth limit is biologically unjustified. The switch between these two communities occurred across bathymetric gradients as small as 10m and as large as 50m in depth. The difference between communities appears to be a loss of shallow specialists and increase in depth-generalist taxa. Therefore, it may be possible to define MCEs by a loss of shallow specialist species. To support a biological definition of mesophotic reefs, we advocate this analytical framework should be applied around the Caribbean and extended into other ocean basins where MCEs are present.

Mesophotic coral ecosystems (MCEs) and temperate mesophotic ecosystems (TMEs) have received increasing research attention during the last decade as many new and improved methods and technologies have become more accessible to explore deeper parts of the ocean. However, large voids in knowledge remain in our scientific understanding, limiting our ability to make scientifically based decisions for conservation and management of these ecosystems. Here, we present a list of key research and con-servation questions to enhance progress in the field. Questions were generated following an initial open call to MCE and TME experts, representing a range of career levels, interests, organizations (including academia, governmental, and nongovernmental), and geographic locations. Questions were refined and grouped into eight broad themes: (1) Distribution, (2) Environmental and Physical Processes, (3) Biodiversity and Community Structure, (4) Ecological Processes, (5) Connectivity, (6) Physiology, (7) Threats, and (8) Management and Policy.

Background: Globally, shallow-water coral reef biodiversity is at risk from a variety of threats, some of which may attenuate with depth. Mesophotic coral ecosystems (MCEs), occurring from 30 to 40 m and deeper in tropical locations, have been subject to a surge of research this century. Though a number of valuable narrative reviews exist, a systematic quantitative synthesis of published MCE studies is lacking. We conducted a systematic review to collate mesophotic research, including studies from the twentieth century to the present. We highlight current biases in research effort, regarding locations and subject matter, and suggest where more attention may be particularly valuable. Following a notable number of studies considering the potential for mesophotic reefs to act as refuges, it is important to know how comprehensive these sources of recruits and organisms capable of moving to shallow water reefs may be. Methods: We search seven sources of bibliographic data with two search strings, as well as personal libraries. Articles were included if they contained species presence data from both shallower and deeper than 30 m depth on tropical coral reefs. Studies were critically appraised based on the number of species identified and balanced sampling effort with depth. Maximum and minimum depths per species were extracted from each study, along with study region and taxon. We quantified the degree of community overlap between shallow tropical reefs (< 30 m) and reefs surveyed at the same locations below 30 m. Proportions of shallow species, across all studied taxa, observed deeper than 30 m were used to generate log odds ratios and passed to a mixed-effects model. Study location and taxon were included as effect modifiers. Funnel plots, regression tests, fail safe numbers, and analysis of a high validity subgroup contributed to sensitivity analyses and tests of bias. Results: Across all studies synthesised we found two-thirds of shallow species were present on mesophotic reefs. Further analysis by taxon and broad locations show that this pattern is influenced geographically and taxonomically. Community overlap was estimated as low as 26% and as high as 97% for some cases. Conclusions: There is clear support for the hypothesis that protecting mesophotic reefs will also help to conserve shallow water species. At the same time, it is important to note that this study does not address mesophotic-specialist communities, or the ecological forces which would permit refuge dynamics. As we limit our analysis to species only present above 30 m it is also possible diversity found exclusively deeper than 30 m warrants protection in its own right. Further research into relatively ignored taxa and geographic regions will help improve the design of protected areas in future.

This is an open access article under the terms of the Creat ive Commo ns Attri bution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Background: Mesophotic coral ecosystems (MCEs) are tropical and sub-tropical reefs between 30 m and potentially >150 m depth, the maximum for photosynthetic hard corals. The definition's upper boundary is ecologically arbitrary. Recently, research has focused on the deep reef refugia hypothesis suggesting MCEs can be protected from shallow-water threats, potentially acting as a local source for re-colonisation of shallow reefs. This led to recent calls to increase their protection. It remains unclear whether the current MCE definition reflects changing biodiversity with depth, and so whether protecting MCEs based on this definition will protect shallow reef species. We ask where shifts in ecological community structure occur across the shallow-mesophotic depth gradient. We consider to what extent MCEs as currently defined protect shallow reef taxa. Research on coral reef depth gradients has a long history. Research relevant to MCEs has been published under a variety of terms. We will use the systematic review framework to collect older data sources, increasing accessibility by depositing the meta-data in an online library for researchers and managers. Methods:A systematic review will be conducted, searching online databases, grey literature and personal libraries of experts. The primary question was formulated after consulting an advisory committee. Inclusion criteria discriminate among studies by sampling depths and community data. Critical appraisal of studies will consider key criteria concerning internal validity. We shall identify where more biodiversity and community-level data are required, determined by whether a meta-analysis is possible. Considering how to structure a meta-analysis once community metric and variability data have been collected will help to advise future data collection. Provided enough data are extracted, we shall conduct a meta-analysis examining changes in species richness, abundance and biomass across the depth gradient. If ecological community level data are present, we shall conduct an additional meta-analysis looking at community turnover with depth.

Light quality is a crucial physical factor driving coral distribution 32 along depth gradients. Currently, a 30 m depth limit, based on SCUBA regulations, 33 separates shallow and deep mesophotic coral ecosystems (MCEs). This definition, 34 however, fails to explicitly accommodate environmental variation. Here, we posit a 35 novel definition for a regional or reef-to-reef outlook of MCEs based on the light vs. 36 coral community-structure relationship. A combination of physical and ecological 37 methods enabled us to clarify the ambiguity in relation to that issue. To characterize 38 coral community structure with respect to the light environment, we conducted wide-39 scale spatial studies at five sites along shallow and MCEs of the Gulf of Eilat/Aqaba 40 (0-100 m depth). Surveys were conducted by Tech-diving and drop-cameras, in 41 addition to one year of light spectral measurements. We quantify two distinct coral 42 assemblages: shallow (<40 m), and MCEs (40-100 m), exhibiting markedly different 43 relationships with light. The depth ranges and morphology of 47 coral genera, was 44 better explained by light than depth, mainly, due to the Photosynthetically Active 45 Radiation (PAR) and Ultra Violet Radiation (1% at 76 m and 36 m, respectively). 46Branching coral species were found mainly at shallower depths i.e., down to 36 m. 47Among the abundant upper mesophotic specialist-corals, Leptoseris glabra, Euphyllia 48 paradivisa and Alveopora spp., were found strictly between 36-76 m depth. The only 49 lower mesophotic-specialist, Leptoseris fragilis, was found deeper than 80 m. We 50 suggest that shallow coral genera are light-limited below a level of 1.25% surface 51 PAR and that the optimal PAR for mesophotic communities is at 7.5%. This study 52 contributes to moving MCEs ecology from a descriptive-phase into identifying key 53 ecological and physiological processes structuring MCE coral communities. 54Moreover, it may serve as a model enabling the description of a coral zonation world-55 wide on the basis of light quality data. 56 3